In the present study we investigated in vivo length changes in the fascicles and tendon of the human gastrocnemius medialis (GM) muscle during walking. The experimental protocol involved real-time ultrasound scanning of the GM muscle, recording of the electrical activity of the muscle, measurement of knee-and ankle-joint rotations, and measurement of ground reaction forces in six men during walking at 3 km h À1 on a treadmill. Fascicular lengths were measured from the sonographs recorded. Musculotendon complex length changes were estimated from anatomical and joint kinematic data. Tendon length changes were obtained combining the musculotendon complex and fascicular length-change data. The fascicles followed a di¡erent length-change pattern from those of the musculotendon complex and tendon throughout the step cycle. Two important features emerged: (i) the muscle contracted near-isometrically in the stance phase, with the fascicles operating at ca. 50 mm; and (ii) the tendon stretched by ca. 7 mm during single support, and recoiled in push-o¡. The behaviour of the muscle in our experiment indicates consumption of minimal metabolic energy for eliciting the contractile forces required to support and displace the body. On the other hand, the spring-like behaviour of the tendon indicates storage and release of elastic-strain energy. Either of the two mechanisms would favour locomotor economy.

Architectural properties of the triceps surae muscles were determined in vivo for six men. The ankle was positioned at 15 degrees dorsiflexion (-15 degrees) and 0, 15, and 30 degrees plantar flexion, with the knee set at 0, 45, and 90 degrees. At each position, longitudinal ultrasonic images of the medial (MG) and lateral (LG) gastrocnemius and soleus (Sol) muscles were obtained while the subject was relaxed (passive) and performed maximal isometric plantar flexion (active), from which fascicle lengths and angles with respect to the aponeuroses were determined. In the passive condition, fascicle lengths changed from 59, 65, and 43 mm (knee, 0 degrees; ankle, -15 degrees) to 32, 41, and 30 mm (knee, 90 degrees ankle, 30 degrees) for MG, LG, and Sol, respectively. Fascicle shortening by contraction was more pronounced at longer fascicle lengths. MG had greatest fascicle angles, ranging from 22 to 67 degrees, and was in a very disadvantageous condition when the knee was flexed at 90 degrees, irrespective of ankle positions. Different lengths and angles of fascicles, and their changes by contraction, might be related to differences in force-producing capabilities of the muscles and elastic characteristics of tendons and aponeuroses.

Muscle force (MF) is linearly related to physiological cross-sectional area (CSA), which is obtained from muscle volume (MV) divided by fibre length. Taking into account the fact that joint torque (TQ) is determined by MF multiplied by the moment arm, the maximal TQ would be a function of MV. This proposition was tested in the present study by investigating the relationship between MV and TQ for elbow flexor (EF) and extensor (EE) muscles of 26 males. The MVs of EF and EE were determined from a series of muscle CSA by magnetic resonance imaging (MRI), and pennation angle (theta) and FL by ultrasonography (US). Maximal isometric TQ was measured at right angle of elbow joint for EF and EE. There was a highly significant correlation between MV and TQ both for EF and EE (r=0.95 and 0.96 respectively) compared with that between muscle CSA and TQ, suggesting the dependence of TQ on MV. Furthermore, prediction equations for MV (MVULT) from muscle thickness (MT) measured by US was developed with reference to MVMRI by the MRI on 26 subjects, and the equations were applied to estimate MV of healthy university students (CON; 160 males) and sports athletes (ATH; 99 males). There were significant linear relationships between MVULT and TQ both for EF (r=0.783) and EE (r=0.695) for all subjects (n=259). The MVULT was significantly higher in ATH (by 32% for EF and 33% for EE, respectively) than in CON. Similarly, significantly greater TQ was observed in ATH (by 35% for EF, 37% for EE, respectively). The theta for EE showed no difference between both groups (17.8 degrees for CON and 17.5 degrees for ATH). On the other hand, the TQ to MV ratio were identical for CON and ATH. The results reveal that the muscle volume of the upper arm is a major determinant of joint torque (TQ), regardless of athletic training.

Muscle-fiber pennation angles were measured in vivo with the use of ultrasonography to investigate the relationship between fiber pennation and muscle size for 32 male subjects (from untrained subjects to highly trained bodybuilders). From the image of a B-mode ultrasonogram, fiber pennation angles and thickness of triceps brachii were determined, the former as angles between echoes from the interspaces of fascicles and from the aponeurosis of long and medial heads of triceps and the latter as the distance between the fat-muscle and muscle-bone interfaces. The pennation angles were in the range of 15 and 53 degrees for the long head and 9 to 26 degrees for the medial head, which were similar to or greater than the published and the present data on human cadavers. Significant differences were observed between normal subjects and bodybuilders in muscle thickness and pennation angles (P < 0.01), and there were significant correlations between muscle thickness and pennation angles for both long (r = 0.884) and medial (r = 0.833) heads of triceps, suggesting that muscle hypertrophy involves an increase in fiber pennation angles.

We have developed a technique to determine fascicle length in human vastus lateralis muscle in vivo by using ultrasonography. When the subjects had the knee fully extended passively from a position of 110 degree flexion (relaxed condition), the fascicle length decreased from 133 to 97 mm on average. During static contractions at 10% of maximal voluntary contraction strength (tensed condition), fascicle shortening was more pronounced (from 126 to 67 mm), especially when the knee was closer to full extension. Similarly, as the knee was extended, the angle of pennation (fascicle angle, defined as the angle between fascicles and aponeurosis) increased (relaxed, from 14 to 18 degrees; tensed, from 14 to 21 degrees), and a greater increase in the pennation angle was observed in the tensed than in the relaxed condition when the knee was close to extension (< 40 degrees). We conclude that there are differences in fascicle lengths and pennation angles when the muscle is in a relaxed and isometrically tensed conditions and that the differences are affected by joint angles, at least at the submaximal contraction level.

Magnetic resonance imaging techniques were used to determine the physiological cross-sectional areas (PCSAs) of the major muscles or muscle groups of the lower leg. For 12 healthy subjects, the boundaries of each muscle or muscle group were digitized from images taken at 1-cm intervals along the length of the leg. Muscle volumes were calculated from the summation of each anatomical CSA (ACSA) and the distance between each section. Muscle length was determined as the distance between the most proximal and distal images in which the muscle was visible. The PCSA of each muscle was calculated as muscle volume times the cosine of the angle of fiber pinnation divided by fiber length, where published fiber length:muscle length ratios were used to estimate fiber lengths. The mean volumes of the major plantarflexors were 489, 245, and 140 cm3 for the soleus and medial (MG) and lateral (LG) heads of the gastrocnemius. The mean PCSA of the soleus was 230 cm2, about three and eight times larger than the MG (68 cm2) and LG (28 cm2), respectively. These PCSA values were eight (soleus), four (MG), and three (LG) times larger than their respective maximum ACSA. The major dorsiflexor, the tibialis anterior (TA), had a muscle volume of 143 cm2, a PCSA of 19 cm2, and an ACSA of 9 cm2. With the exception of the soleus, the mean fiber length of all subjects was closely related to muscle volume across muscles. The soleus fibers were unusually short relative to the muscle volume, thus potentiating its force potential.(ABSTRACT TRUNCATED AT 250 WORDS)

These results indicate that the jump performance gains after plyometric training are attributed to changes in the mechanical properties of muscle-tendon complex, rather than to the muscle activation strategies.

Reported specific tension measurements for human skeletal muscle vary widely. This variability could be due, at least in part, to the determination of the physiological cross-sectional area (PCSA) of the muscles. In the present study, serial magnetic resonance images were taken every 10 mm along the lower leg of 8 male subjects to calculate the volume and subsequently the PCSAs of the individual muscles producing plantar flexor and dorsiflexor torques. Maximum plantar flexor and dorsiflexor voluntary isometric torques were determined at ankle joint angles of 90, 100, 110, and 120 degrees. Peak tendon force estimated from torque and moment arm measurements was more than fourfold higher in the plantar flexors (3,623 +/- 136 N) than in the dorsiflexors (832 +/- 19 N). PCSAs were about eight- and threefold higher than the anatomic cross-sectional areas at the level of maximum girth of the calf for the plantar flexor and dorsiflexor groups, respectively. Mean muscle volume and PCSA were 4.6 and 12 times larger in the plantar flexors compared with the dorsiflexors, respectively. The PCSAs of both plantar flexors (r = 0.92) and dorsiflexors (r = 0.80) were highly correlated with the tendon tension of the respective muscle groups. The maximum specific tension was more than twofold higher in dorsiflexors than in plantar flexors. These data suggest that factors other than PCSA contribute to the force output potential of ankle plantar flexors and dorsiflexors in humans.